Species concept and speciation.pdf - Free download as PDF File (.pdf), Text File (.txt) or read online for free. 0000016101 00000 n Among which 2, 35,000 are angiosperms, 600 gymnopserms and 10,000 pteridophytes. (iii) Common occurrence of interspecific hybridization between species of flowering plants. Intraspecific Categories. Share Your PPT File. It is the largest group or population which is well defined, traditionally as Plant and Animal. Disclaimer Copyright, Share Your Knowledge Sometimes the specific epithet carries its own name, e.g., in Pyrus malus. government site. 8 0 obj <> (b) Species is easily recognized kind of organisms, and in macroscopic plants and animals their recognition should rest on simple gross observation (May be with hand lens only). The cookie is used to store the user consent for the cookies in the category "Analytics". roylei (for Royale); hookeri (for Hooker) etc. Indeed, several species concepts, based on more or less compatible criteria, have previously been proposed (reviewed in De Queiroz 2007). 1 0 obj The names of subspecies are trinomial, the third epithet being of subspecies. Bremer and Wanntorp (1978) recognised six major groups based on a cladistic reinterpretation of Takhtajans (1969) phyletic classification. The need to use more than one spe-, species concept. Species are changing in the sense that each species has its own characteristics and there is no complete identity between any two species. Give an example. Family and Higher Categories. 0000016079 00000 n as species. Then species was defined on a priori basis and regarded as unchanging and fixed. According to it the first step in speciation is reproductive isolation brought about by physical separation or geographical separation of population (allopatric population). (iv) Inapplicability of the concept to asexual species. 0% 0% found this document not useful, . Love (1954) accorded species status to morphologically indistinguishable cytotypes. Accessibility 0000002630 00000 n the questions arises as whether species might not better be regarded as individuals rather than classes. A book in, including 161 families and 58000 species was his main and most, Years later, in his theory of evolution with Wallace (, cies could be produced rapidly if the conditions were appropriate, unchanged for a long time. The problem in this evolutionary, its between species, especially those which experiencing regular, arena of action of basic micro evolutionary forces, such as gene, ). Zh Obshch Biol. Morphologic species or Morphospecies concept is also called as classical phenetic species concept or Linnaean or classical species concept. Mayr (1942, 47, 52) proposed the rule that we should name populations and not species, though an individual can be chosen as the nomenclatural type. 0000003651 00000 n 8H?de0Q%H! 8On-`q Earlier the taxonomists were not knowing that the individual species possess certain degree of capacity for variation. In this paper some of species concepts are discussed starting from the typological species concepts, to the phylogenetic concept. In Angiosperms Orders are not well defined except for a few such as Caryophyllales. Biological species concept the most eminent and momentous in the taxonomy field, adopting a broader concept gave a new definition of species. 2015 Nov;324(7):588-604. doi: 10.1002/jez.b.22636. Answers to the questions of what a species is, in what ways species really exist (if in fact . It results in hybrid formation, pollination, dispersal, population size etc. Sub-sections are good and helpful when a genus has a large number of Species (Infra generic), e.g., Senecio (Asteraceae) has more than 1,000 species. HN@{UUAmI^^MmDFVV=a%'WCfHDYZsRSc 0000005595 00000 n (x) To avoid using the names of little known or very restricted localities unless the species is quite local. The site is secure. The name of an infraspecific taxon is a combination of the name of a species and an infraspecies epithet connected by a term denoting its rank. 0000014538 00000 n A population of one or several biotypes, forming more or less distinct local facies of a species. Problems with the biological species concept The Biological Species Concept (BSC) is not perfect:-Does not apply to asexual organisms, which include many microbes and some plants-Difficult to apply or test-Hybridization: many species (especially plants) can interbreed at least a little with other species That is why scientists also use many other species concepts! It starts with the great Philosopher Plato who proposed concept of eidos or species and believed that all objects are shadows of the eidos. 0000006185 00000 n Functional cookies help to perform certain functionalities like sharing the content of the website on social media platforms, collect feedbacks, and other third-party features. There are many isolating mechanisms which operate to prevent or reduce the gene flow between populations: It depends upon some aspects of plants themselves such as lock and key pollination mechanism, difference in flowering time of the two populations etc., e.g., Salvia apiana and S. mellifera are two species growing in different zones but show resemblances also. He divided plants into, two major groups, non-vascular and vascular plants. After many years (Aristotle) species was defined on an a priori basis and was regarded as fixed and unchanging. Variation is a deviation in structural, functional or developmental characters of an organism from its parents, from others in the same population. 0000006672 00000 n 0000010128 00000 n Broadly; 5 different concepts are given: It believes that Nature produces individuals and nothing more. Variant is an individual or a group of individuals within a population that is definiable and recognizable. 0000011954 00000 n (9) To give feminine forms to all personal generic names, whether they commemorate a man or woman. Species concept and speciation. (iv) To avoid those formed if two or more hyphenated words. A CITESlisted species might relate to - multiple species on the IUCN Red List (or vice versa), likewise a CITES-listed subspecies . Various attempts have been made to define a species. Populations are therefore dynamic and show variations with in the population and variation between the populations of the same species. 0000004562 00000 n (e) Environment of the individuals must not be static. Compilospecies is one which is morphologically distinct (Taxonomic species concept) but not reproductively isolated (single species according to biological concept), e.g., Salvia mellifera and S. apiana which are morphologically distinct but reproductively not isolated. 0000009053 00000 n !bw1TFK 0'0 endstream endobj 172 0 obj 765 endobj 173 0 obj << /Filter /FlateDecode /Length 172 0 R >> stream This concept is useful and meaningful even for those plants where hybridization is common (e.g., Quercus). It does not store any personal data. It is of three fundamental types, i.e., developmental, environmentally induced and genetic variation. (2) To avoid names not readily adaptable to the Latin language. The cookies is used to store the user consent for the cookies in the category "Necessary". Box 80200, Jeddah 21589, Saudi Arabia, Dening and recognizing a species has been a controversial issue for a long time. This monotypic or static concept prevailed till the 19th century. stream Mutation may be as small as substitution of single nucleotide pair in the DNA molecule or as big as chromosomal aberration (structure or number). Species are man's own creations and have no actual existence in nature. (2) Biological species: (Genetic species): (4) Successional species: (Paleospecies): (5) Biosystematic species: (Ecospecies, Coenospecies): Combined sexually reproducing populations iniparental groups and phyletic lineages. Analytical cookies are used to understand how visitors interact with the website. Linnaeus used only one infraspecific taxon, the variety. Bethesda, MD 20894, Web Policies 0000005970 00000 n (ii) Typological Species Concept or Taxonomic species concept: According to it species is A very natural group of organisms hence a natural taxon in classification has an invariant generalized or idealised pattern shared by all members of the group. The species, as we know, is the fundamental unit of taxonomic hierarchy. In biological phenomenon biosystematics concept is [] The main use of species in taxonomy and derivative sciences is to order and retrieve information on individual specimens in collections or data banks. If the two populations grown side by side, may cross and produce hybrids that cannot tolerate the lead concentration on one side or cannot compete with the normal plants of the species on the other side. H\TM6(|+"%mdEbk,lII36:hHyffSm^+m[, endobj Krukeberg (1969) stressed that the bio-species also differ in their ecological contexts. G13;PuS+=P:J Zh Obshch Biol. w%@"@ `%B&.. A%La`@&1~eX8t1~aX q! Vj@.n* sYP]!5ArCc:tVMW- 0000006851 00000 n Species: Concept, Types and Intraspecific Categories. In the accepted system of nomenclature every individual plant is treated as belonging to a number of taxa of consecutively subordinate ranks; each with subcategories or taxa of consecutively as ascending rank e.g., an individual plant of swamp rose would belong to the consecutively higher ranks of the species. According to this concept, "Nature produces individuals and nothing more. on the origins of phenotypic similarity within species. P.367-368)]. When a given species concept is favored in a given conditions, that does not mean it could be universally applicable. Comparing, with BCS this concept is applicable on both sexual and allopatric. Thus, it would clarify some particular degree of, Practically, the phenetic concept measures as many characters, as possible in as many organisms as possible, and then identies, phenetic clusters by multivariate statistics. (viii) To avoid those which have been used before in any closely allied genus. %PDF-1.3 % Disclaimer, National Library of Medicine Necessary cookies are absolutely essential for the website to function properly. It is thus a genodeme orrelatively minor genetic variant occurring mixed with other such distinct variants. The problem comes in knowing the genetic difference between populations. It can be applied to sexual and asexual organisms and it is, depend on expert opinion for key traits. This concept avoids the difficulties with determining actual or potential levels of interbreeding as gene flow and allows some degree of inter-specific hybridization. This unified species concept has a num- ber of consequences for taxonomy, including the need to acknowledge that undiffer- entiated and undiagnosable lineages are species, that species can fuse . Those complex of species which cannot be sorted out taxonomically from a population and in which there is hope of eventual resolution are called species aggregate or Microspecies. The taxa are referred to particular categories, e.g., species or variety available in the hierarchy. Each concept has its own advantages, but also its own approximations of the biological truth, so it now appears evident to integrate multiple criteria and go towards a unified species concept (De Queiroz 2005). 0000002951 00000 n [Some notes on relation between taxon and character in taxonomy (regarding the paper of A.A. Stekol'nikov "A problem of truth", Zhurnal Obshchei Biologii. In his, and sexual characters, Linnaeus described briey and systemati-. (Table 1). (ii) To avoid epithets which are very long and difficult to pronounce in Latin. Taxonomy of diatoms. W.z,\. Unable to load your collection due to an error, Unable to load your delegates due to an error. Family and Higher Categories of Species Concept: This cookie is set by GDPR Cookie Consent plugin. To determine the vari-, ation and the limitation between species, many concepts have been proposed. description of species as ecologically or genetically meaningful entities with a shared phylogenetic heritage. Recombination is a reassortment of chromosomes, crossover segments etc. We also use third-party cookies that help us analyze and understand how you use this website. Separation results commonly from shrinkage of the original range of a species so that the populations become physically isolated. Similar but hybrids between them are of reduced viability. In addition, if a study is concerned on the simi-, larity of a group of plant with enough information of characters, (morphology, anatomy, cytology) with no need to a lineage, a mor-, phological concept could be adapted. xJ bD+#,.HRmRmDE|pKK 2 + @ @5 lmm[[7o{c! PMC 0000011065 00000 n According to this concept, species is a spatio temporal lineage of populations that evolves separately from other lineages and has its own ecological niche. Variety is the term applicable to diversity, variant population, variability, character variation etc. Due to first step in differentiation ecotypes are formed. 0000008429 00000 n ! LT4W&>Tn#?1_MVj$Tw&m@5ihJ% z4Z"I%-iNIU+Gc/VFj9N[" v*()4dw6/p/ >s( XOmm gI++VEo|edNr)!29kq[z]_COt:_A>CkxH|6( hm hv]jZN `ne_Y~n.~OEswQ>e|cF/QP In 1872, Ferdinand Cohn demonstrated that bacteria could be divided into genera and species using the paradigm proposed for plants and animals by the father of modern taxonomy, Carl Linnaeus. 0000026518 00000 n 2012;61:79-209. doi: 10.1016/B978-0-12-387787-1.00010-6. Diatoms belong to a large group called the heterokonts, which include both autotrophs such as golden algae and kelp; and heterotrophs such as water moulds. Since then, many modern species concepts were developed. Share Your Word File 2017 Production and hosting by Elsevier B.V. on behalf of King Saud University. To approach a satisfactory and acceptable classi-. 2004 Jul-Aug;65(4):334-66. The genetic species concept assumes that the biological factors of gene flow and reproductive isolation are operative but the way to define species is by a measure of the genetic differences or distance among population. Concept of Species 2. All specific epithets should begin with small letter of English. They believed that taxonomy means that the description of a single plant is the description of the species to which it belonged. 0000012736 00000 n Combining the viewpoints of cladistics and evolutionary systematics, the writer proposes six rules for character analysis which will provide the basis for analyzing evolutionary history. 0000032211 00000 n endstream What is a trophic hormone? Character trees from transcriptome data: Origin and individuation of morphological characters and the so-called "species signal". According to Grant (1981) Spatial or geographic isolation exists between any allopatric species whose respective geographical areas are separated by gaps greater than the normal radius of dispersal of their pollens or seed, i.e., Liriodendron tulipifera of Eastern United State and L. chinense of South eastern Asia, which cannot breed under natural conditions but have been crossed artificially. Davis (1978) called them Building bricks in Biological classification. ADVERTISEMENTS: In this article we will discuss about Species:- 1. (3) Not to make names which are very long and difficult to pronounce in Latin. 0000015332 00000 n x On the other hand cladistic or phylogenetic basically view evo-. Van Valen (1976) defined it as A species is a lineage occupying an adaptive zone minimally different from that of any other lineage in its range and which evolves separately from all lineages outside its range.. After the knowledge of a number of organisms, people started facing difficulty as there are species which belong to different genera. He felt that the numerical phenetic species concept has more problems, particularly on deciding which phenetic unit is the one to be called species. Simpson (1961) also suggested that niche included multidimensional relationship of a taxon to its environment and not just its microgeographic situation. Save Save Species Concepts.pdf For Later. Thus groups of related plants, which are distinct at the level of biological species do not inter-, breed when growing in the same area in nature. To achieve the ranking in Taxonomy a hierarchy of categories is suggested. genetic species; polytypic species; species limits. Taxon means a cluster of individuals grouped together based on the sharing of features in common. Linnaeus in Philosophia Botanica (1751) used 3 characters for generic concepts, i.e. But now subspecies and varieties are also available. Name the types of nitrogenous bases present in the RNA. Species is the fundamental unit of it. ~dS%U"c+?%m>*0?dc j`wgyzvo koK%5i]?KV~\k}-M=cTRvd! ~{rYP*vf4o*J ra- akjyMR*=P^,dJ:S~n6 73MmJ6!su@FSQaBg|*9*UjFZV#Lg0T^U f ub#V+ This website includes study notes, research papers, essays, articles and other allied information submitted by visitors like YOU. The species reproduce only vegetative propagation, e.g., Phragmites. Vaughan (1905) termed a collection of paleospecies in a monophyletic succession as genes. Asters for the genus Aster. (ii) Real extent of gene flow among population. Now-a-days only three of the five suggested by ICBN are in use. 0000003081 00000 n Species Concepts Jody Hey Introduction In a general and nontechnical sense, species are kinds of organisms. Largely, the importance of species stems from its significance in systematics, which is a science all the branches of biology rely on . Agrostis tenuis, a temperate grass which also grown on lead concentrated soil, and becomes lead tolerant. % C A_{ aL endobj Donoghue (1985) and Mishler (1985) called it as phylogenetic species concept. 7 0 obj Speciation may take place abruptly also by development of polyploids and mutations Futuyma (1983) suggested that Speciation more thoroughly a wash in unfolded and often contradictory speculation than any other single topic in evolutionary theory. There are many others (ex. Diversity can be seen in various types also, e.g., in Rosaceae at least 9 fruit types are found. These cookies track visitors across websites and collect information to provide customized ads. : (a) Natural character (complete description of plant). The specific name may be pronoun, adjective etc. (7) Not to use a name similar to or derived from the epithet in the name of one of the species of the genus. Species concepts do not only dene what a species is, but by dening what a species is, they also clarify what speciation, is. These cookies will be stored in your browser only with your consent. 0000031580 00000 n In this article we will discuss about:- 1. Tournefort (1700) attempted to make informal groups of genera with some similar characters and tried to describe them. What are the characters Mendel selected for his experiments on pea plant? Genus is next to species as far as taxonomic hierarchy is considered. etc. This situation creates a problem given that species are used as basic units of comparison in diverse types of stud- ies. A variety may be quadrinomial as the fourth epithet will be for varieties but in case there is no subspecies, it will be trinomial, e.g., Brassica oleracea var. 0000012714 00000 n According to Stace (1989) the two species are separated by the ecological and mechanical differences. HHS Vulnerability Disclosure, Help Types 4. What do you mean by permeability of membrane? Therefore, it can combine. endobj The, theory of phenetic species concept can be opposed on the bases, of that, to a specic degree there is a resemblance between any, two objects in the universe. 0000004961 00000 n Light microscopy of several species of living freshwater diatoms. lution as an ordered and divergent transformation of characters. The .gov means its official. Secondly, it is impractical in instances of allo-, Both isolating species concept and recognition species concept, can be part of biological concept or genetic concept because both, see the species as a eld for gene recombination. In the middle of the twenty century, exactly in 1920s, a new, science was born. 0000005801 00000 n The cookie is used to store the user consent for the cookies in the category "Other. present in taxonomic world. Although this theory is so simple and obvi-, ous, it has mainly two disadvantages. With this large number of concepts it. stream The term family was first used by A.L. On the other hand, species are unchanging and each one conserves a set of ancestral characters which allocates its taxonomic position, and this is the reason why the taxonomic system reflects evolutionary . and which evolves separately from all lineages outside its range. (vi) To avoid those which express a character common to all or nearly the species of a genus. It includes experimental taxonomic studies also. 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In his major work, ) he stated: No matter what variations occur, in the individuals or the species, if they spring from the seed of one, and the same plant, they are accidental variations and not such as, distinguish a species permanently; one species never springs from, dene species as groups of plant truly breed within their limits. The generic name should be accompanied with a specific epithet following the binomial system of Linnaeus. modern science. History of Species Concept 2. Thus the cohesion concept is similar to, lutionary species concept in a way that a population genetic stress. xUH}G-MvH 0Y D+b C2 mFyv)Ox+x3nGB`.^@HXmH0- i[W1?PB67hPm{ U Evolutionary concept rejects the nominalistic concept of species. FOIA FPUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUa _AUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUUU=8 uC ]!J

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